Lecture 2 The plasma membrane
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1 Lecture 2 The plasma membrane
2 Plasma membranes: 1. Lipid composition of cell membranes 2. Singer and Nicolson fluid mosaic model 3. Proteins associated with the plasma membrane can be integral or peripheral 4. Functions of membrane proteins 5. Microdomain organization of the plasma membrane 6. FRAP: A method for measuring diffusibility in the membrane
3 Cell membranes surround all cellular compartments Functions 1. compartmentalization 2. scaffolding for biochemical activities (e.g. energy production) 3. selective transport 4. reception of stimuli from the environment 5. interactions with neighboring cells (e.g. adhesion)
4 Composition of cell membranes Observation by electron microscopy (1960s) revealed an electron-dense bilayer separated by a clear layer (lipids) 2
5 Structure of phospholipids in cell membranes hydrophillic head choline phosphate glycerol non-polar hydrophobic tail images from Nature Education website
6 Cell membranes are bilayers of amphipathic lipids
7 The chemical nature of membrane lipids allow them to self-assemble into membranes
8 The chemical nature of membrane lipids allow them to self-assemble into membranes Detergents dissociate plasma membranes
9 The cell membrane is a dynamic structure that is constantly renewed ER Membrane is made in the ER and added to the plasma membrane by vesicle fusion. When vesicles fuse to the plasma membrane their lipids are incorporated into it. Endocytosis brings membrane back into the cell.
10 The chemical synapse: An extremely dynamic membrane region
11 Intracellular membranes Membranes that form organelles have the same basic plasma membrane structure. However, the specific composition of lipids and proteins differs significantly between organelles and from cell to cell
12 Cell membranes contain many chemically distinct kinds of lipids with different properties
13 Different membranes in a cell contain different complements of lipids van Meer G, Voelker DR, Feigenson GW (2008). Membrane lipids: where they are and how they behave. Nat Rev Mol Cell Biol. 9(2):
14 How do different lipids end up in different membranes? ER Selective transport: Lipids made in the ER or incorporated from the outside are trafficked to different destinations in a cell.
15 How do different lipids end up in different membranes? PI3kinase Lipids can be phosphorylated or dephosphorylated to endow their head groups with different chemical properties
16 How do different lipids end up in different membranes? Lipids can be cleaved by lipases
17 How do different lipids end up in different membranes? Flippases, floppases and scramblases are enzymes that use energy from ATP to exchange lipids from one leaflet to another Clark, MR (2011). Flippin' lipids. Nature Immunology 12:
18 How do different lipids end up in different intracellular membranes? Newly formed lipids are transported to specific locations Lipids can be covalently modified; for example, they can be phosphorylated, dephosphorylated or cleaved Lipids can be flipped from one leaflet to another PI3kinas
19 Lipids and proteins in the membrane can diffuse laterally-- fluid mosaic Singer and Nicolson model
20 Singer and Nicolson (1972) Science
21 Singer and Nicolson (1972) Science
22 Temperature affects the fluidity of the lipid bilayer Above the transition temperature, the molecules of the lipid bilayer are free to move. Below the transition temperature, the movement of molecules is reduced and restricted.
23 Membranes contain many associated proteins and carbohydrates Singer and Nicolson model 1972
24 Proteins can be associated with membranes in several different ways Transmembrane proteins Peripheral proteins Lipid - anchored proteins
25 Integral membrane proteins can be recognized from their amino acid sequences Transmembrane domains are often alpha helices consisting of hydrophobic amino acid residues Regions of hydrophobicity can be visualize in a Kyte-Doolittle plot
26 Proteins can be covalently conjugated to lipids Lipid modifications regulate cell signaling by targeting intracellular proteins to cell membrane 16
27 An experimental approach for deducing the nature of protein association with plasma membrane How are proteins associated with red blood cell membranes?
28 Studying proteins of the plasma membrane Solubilize the membrane and run proteins on a gel
29 Studying proteins of the plasma membrane Add trypsin to digest proteins on outside of cell
30 Studying proteins of the plasma membrane Permeabilize and add trypsin to digest proteins on inside and outside of cell
31 Studying proteins of the plasma membrane
32 Membranes contain many associated proteins and carbohydrates Singer and Nicolson model 1972
33 Functions of Membrane Proteins image from Nature Education website
34 Functions of Plasma Membrane Proteins - Receptors Signal Receptors recognize specific signals from the outside to promote transcription, proliferation, survival, death, changes in cell shape, migration, etc
35 Functions of Plasma Membrane Proteins - Attachment Receptors on the surface of cells can be used to attach to the extracellular matrix or to other cells
36 Functions of Plasma Membrane Proteins - Transport
37 The plasma membrane is organized into domains Distribution of proteins within the plasma membrane can change to accommodate the functional demands of the cell
38 Proteins move within the lipid bilayer Depending on the cell type, the specific protein and environmental conditions, integral membrane proteins can exhibit several types of mobility.
39 The structure of the plasma membrane consists of discrete microdomains that segregate functions into subcompartments yeast cell microdomains Both lipids and proteins are segregated into microdomains
40 Cholesterol- and sphingolipid-rich lipid rafts are microdomains that are thought to concentrate signaling proteins From the inner life of the cell
41 Surveying membrane localization in yeast Tagged Proteins Lipid reporters Most, if not all, proteins localize in non-uniform patches of cell membranes Patchwork organization of the yeast plasma membrane into numerous coexisting domains. Spira F, Mueller NS, Beck G, von Olshausen P, Beig J, Wedlich-Söldner R. Nat Cell Biol Apr 29;14(6):640-8.
42 What factors regulate protein diffusion in the endoplasmic reticulum? Is diffusibility affected by folding status? Is diffusibility affected by sugar modifications? Nehls et al. (2000). Dynamics and retention of misfolded proteins in native ER membranes. Nature Cell Biology 2:288.
43 FRAP-Fluorescence Recovery After Photobleaching A method for measuring protein movement within plasma membranes
44 FRAP-Fluorescence Recovery After Photobleaching A technique that can be used to determine the diffusibility of proteins in the membrane
45 Figure 3: How quickly do proteins move around in the plasma membrane? Patchwork organization of the yeast plasma membrane into numerous coexisting domains. Spira, F., Mueller, N.S., Beck,G.S., von Olshausen, P., Beig, J. and R. Wedlich-Söldner (2012). Nature Cell Biology 14,
46 What factors regulate protein diffusion in the endoplasmic reticulum? Is diffusibility affected by folding status? Is diffusibility affected by sugar modifications? Nehls et al. (2000). Dynamics and retention of misfolded proteins in native ER membranes. Nature Cell Biology 2:288.
47 What factors regulate protein diffusion in the endoplasmic reticulum? VSVG-GFP: An integral membrane protein in the ER that is fused to GFP and expressed in COS cells
48 Is diffusibility affected by folding status? VSVG folds properly at 32ºC, but is misfolded at 40ºC Nehls et al. (2000). Dynamics and retention of misfolded proteins in native ER membranes. Nature Cell Biology 2:288.
49 Is diffusibility affected by sugar modifications? Tunicamycin is a drug that prevents sugar synthesis Nehls et al. (2000). Dynamics and retention of misfolded proteins in native ER membranes. Nature Cell Biology 2:288.
50 What factors regulate protein diffusion in the endoplasmic reticulum? The diffusibility of VSVG in the ER is affected by sugars but not by folding. Nehls et al. (2000). Dynamics and retention of misfolded proteins in native ER membranes. Nature Cell Biology 2:288.
51 Plasma membrane summary: 1. Cell membranes are bilayers of amphipathic lipids 2. The lipid composition of membranes varies 3. Membrane fluidity is regulated by temperature 5. Three types of membrane proteins: 1. Integral, 2. Peripheral, 3. Lipid-anchored 4. Membrane proteins function in signaling, adhesion, and transport 5. The membrane is organized into microdomains 6. Techniques for studying membrane proteins: proteolysis/protection assay, FRAP
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